J. Nannoplankton Res. 31 (1), 2010, pp.39-56 © 2010 International Nannoplankton Association, Inc.
ISSN 1210-8049 Printed by The Sheridan Press, Hanover, PA, USA

39

Calcareous nannofossils from the boreal Upper CampanianMaastrichtian chalk of Denmark
Nicolas Thibault
Department of Geography & Geology, University of Copenhagen, Øster Voldgade 10, DK-1350, Copenhagen K, Denmark; nt@geo.ku.dk

Manuscript received 2nd September, 2009; revised manuscript accepted 18th April, 2010

Abstract Boreal calcareous nannofossil assemblages have been documented from three sections in Denmark, fromthe Upper Campanian to Upper Maastrichtian (nannofossil zones UC16a BP to UC20dBP): the Stevns-1 borehole, next
to the Cretaceous/Palaeogene boundary section of Stevns Klint, eastern Sjælland, the Rørdal-1 borehole and the Rørdal
quarry section, North Jylland. The moderately-preserved nannofossil assemblages consist of a total of 138 species, comprising several rarely-documented species, several species that were recently described from lower latitudes, two new
species of heterococcolith (Prediscosphaera borealensis, Tranolithus stemmerikii), and one new combination (Staurolithites dicandidula).

Keywords Calcareous nannofossils, taxonomy, Campanian, Maastrichtian, boreal, Denmark
1. Introduction
In 2005, the Cretaceous Research Centre (CRC) was established jointly at the Geocenter Copenhagen by the Department of Geography and Geology, University of
Copenhagen, and the Geological Survey of Denmark andGreenland (GEUS). CRC projects aim to study the Earth
System in a Greenhouse World, with special emphasis on
the Upper Cretaceous to Danian chalks of NW Europe. Oneof the main CRC projects was a drilling campaign at Stevns
Klint in Sjælland (eastern Denmark), where two coreholes(Stevns-1 and Stevns-2) were drilled and logged from near
the base of the Lower Danian Bryozoan Limestone, down
through the upper 350m and 450m, respectively, of the
thick Upper Cretaceous Chalk Group (Stemmerik et al.,2006).
The investigation of calcareous nannofossil assemblages from the chalk of the Stevns-1 (Sjælland) andRørdal-1 coreholes, and the Rørdal quarry section (northern Jylland; Figure 1) allowed for the testing of nannofossil biozonation schemes (Sheldon, 2008), and will be used
to document the palaeoecological changes throughout thelatest Cretaceous of Denmark. The sediments are moderately preserved and yield a diverse assemblage, including
two new species, and several species recently describedfrom Tanzania (Lees, 2007) that have not been recorded
previously from boreal latitudes. The aim of this study is toprovide a catalogue of the species encountered in the Danish Upper Campanian–Upper Maastrichtian Chalk Group,
to serve as a guide for future studies in the area, and tocomplement the atlas presented in Burnett et al. (1998).
2. Material
The investigated succession belongs to the upper part of
the Upper Cretaceous Chalk Group (Surlyk et al., 2003).
The Stevns-1 shallow corehole was drilled approximately
2km east of Sigerslev, immediately adjacent to the coastal
cliff of Stevns Klint, in eastern Sjælland (Denmark; Fig-

Figure 1: Late Cretaceous palaeogeography of the Danish area, showing
locations of the Stevns-1 corehole and Rørdal

ure 1). The drilled succession was deposited on the easternend of the Ringkøbing-Fyn High. The Stevns-1 corehole
recovered 456m of sediment from the Lower Danian down
into an Upper Campanian chalk-marl succession with near
100% recovery. The nannofossil biostratigraphy of the core
was near-complete (Sheldon, 2008). Thus, to date, Stevns1 represents the first complete onshore sequence throughthe Upper Campanian–Maastrichtian chalk of NW Europe(compare with Surlyk & Birkelund, 1977; Bailey et al.,
1983; Schulz et al., 1983; Villain, 1988; Schönfeld et al.,
1996; Mutterlose et al., 1998; Surlyk & Håkansson, 1999)
and one of the most expanded Maastrichtian sections,worldwide, in the published literature (compare with sections in von der Borch et al., 1974; Tucholke et al., 1979;

40

Thibault

Hay et al., 1984; de Graciansky et al., 1985; Heath et al.,
1985; Monechi & Thierstein, 1985; Huber, 1992; Mutterlose et al., 1998; Li & Keller, 1998; Li et al., 1999;
Bralower et al., 2002; Howe et al., 2003; Erbacher et al.,
2004; Zachos et al., 2004; Huber et al., 2008).
The Rørdal quarry is situated in the town of Aalborg innorthern Jylland. It is located in the inverted Aalborg
Graben, which forms part of the NW-SE-orientated Sorgenfrei-Tornquist Zone, a region of the Danish Basin characterised by Late Cretaceous and Paleogene structural
inversion (EUGENO-S Working Group, 1988). The quarry
exposes up to 40m of Upper Maastrichtian chalk, including
a c.10m-thick, cyclic chalk-marl unit at the top. Sediments
are highly bioturbated and exhibit trace fossils, such as
Thallassinoides, Zoophycos and Chondrites. The Rørdal-1
corehole was drilled in 1967 at a location immediately adjacent to the quarry wall. It penetrated about 100m ofUpper Campanian-lowermost Upper Maastrichtian chalk,with a few clay laminae and thin marl layers occuring in
the lowermost and uppermost parts of the core (Stenestad,2005). Only chalk samples were examined during thisanalysis. Locality and age details for the samples mentioned here are given in Table 1.
The UCBP (Boreal Province) part of the global nannofossil biozonation scheme of Burnett et al. (1998) was used
to provide a nannofossil biostratigraphy for these cores.
Biostratigraphic information about Stevns-1 is presentedin Sheldon (2008). Full biostratigraphic information about
Rørdal-1 and the Rørdal quarry section will be provided ina separate publication (Thibault et al., in prep.). Preservation of the investigated samples is moderate.
Species richness varies between 40 and 80 species in
the material investigated. This corresponds to 27 to 53%
of the global compilation of nannofossil species for theCampanian–Maastrichtian interval (between 135 and 150:
Bown et al., 2004). This global estimation needs revision,
because a number of new species from this interval havebeen recently described (Lees & Bown, 2005; Lees, 2007).
3. Methods
The species documented here are based on light-microscope (LM) observations only. Nannofossil smear-slides
were prepared following the standard procedure (Bown &Young, 1998), and examination of nannofloras was performed using a Leica DMRP transmitting light microscopeat 1600x magnification. Digital images were captured witha Leica DFC320 digital camera at 512 x 512 pixels, using
UTHSCSA ImageTool 3.0 freeware (available at
http://ddsdx.uthscsa.edu/dig/itdesc.html). All images weretaken in either cross-polarised light (XPL), or in brightfield (BF), the latter being indicated on the plates. Morphometric measurements were performed using UTHSCSA
ImageTool 3.0, a digital-image processing program. At amicroscope magnification of 1600x, the pixel scale is 30

Table 1 (right): Sample locations and biostratigraphy

SampleDepth (m)Age

BiozoneStevns-1 corehole (S1)
1475
13.28
1474
14.29
1473
15.37
1471
17.25
1469
19.00
1467
21.39
1466
22.10
1463
24.42
1462
26.62
1447
40.24
1442
45.95
1432
55.90
1422
64.80
1411
75.00
8180
90.34
1390
95.53
8157
97.69
5623
99
1386
99.43
8147
101.74
5260
102.7
5256
102.87
5253
103
5235
103.721380
104.91
5209
105.3
5620
106.371370
114.81
1364
120.351353
130.731337
145.251310
170.501305
175.22
1300
179.791287
191.241283
194.841272
205.381262
214.601240
234.891229
244.951207
265.491203
269.11
1186
284.81
1175
295.041170
299.631164
305.297398
310.101153
315.417370
320.531141
325.771131
335.061119
345.851114
350.721109
355.341087
375.861083
379.61
1066
395.401055
405.771044
415.781034
425.241029
429.717086
430.257084
430.75
7081
431.5
7076
432.887074
433.577072
433.977070
434.695028
434.921017
441.131011
446.671007
450.481004
453.29Upper MaastrichtianUpper MaastrichtianUpper MaastrichtianUpper MaastrichtianUpper MaastrichtianUpper MaastrichtianUpper MaastrichtianUpper MaastrichtianUpper MaastrichtianUpper MaastrichtianUpper MaastrichtianUpper MaastrichtianUpper MaastrichtianUpper MaastrichtianUpper MaastrichtianUpper MaastrichtianUpper MaastrichtianUpper MaastrichtianUpper MaastrichtianUpper MaastrichtianUpper MaastrichtianUpper MaastrichtianUpper MaastrichtianUpper MaastrichtianUpper MaastrichtianUpper MaastrichtianUpper MaastrichtianUpper MaastrichtianUpper MaastrichtianUpper MaastrichtianUpper MaastrichtianUpper Maastrichtian
Lower Maastrichtian
Lower Maastrichtian
Lower Maastrichtian
Lower Maastrichtian
Lower Maastrichtian
Lower Maastrichtian
Lower Maastrichtian
Lower Maastrichtian
Lower Maastrichtian
Lower MaastrichtianCampanian/MaastrichtianCampanian/MaastrichtianCampanian/MaastrichtianCampanian/MaastrichtianCampanian/MaastrichtianCampanian/MaastrichtianCampanian/MaastrichtianCampanian/MaastrichtianCampanian/MaastrichtianCampanian/MaastrichtianCampanian/MaastrichtianUpper CampanianUpper CampanianUpper CampanianUpper CampanianUpper CampanianUpper CampanianUpper CampanianUpper CampanianUpper CampanianUpper CampanianUpper CampanianUpper CampanianUpper CampanianUpper CampanianUpper CampanianUpper CampanianUpper CampanianUpper CampanianUpper CampanianUpper Campanian

UC20dBP
UC20dBP
UC20dBP
UC20dBP
UC20dBP
UC20dBP
UC20dBP
UC20dBP
UC20dBP
UC20dBP
UC20dBP
UC20dBP
UC20dBP
UC20b-cBP
UC20b-cBP
UC20b-cBP
UC20b-cBP
UC20b-cBP
UC20b-cBP
UC20b-cBP
UC20b-cBP
UC20b-cBP
UC20b-cBP
UC20b-cBP
UC20b-cBP
UC20b-cBP
UC20b-cBP
UC20b-cBP
UC20b-cBP
UC20b-cBP
UC20aBP
UC20aBP
UC19
UC19
UC19
UC19
UC19
UC19
UC19
UC19
UC17-18
UC17-18
UC16dBP
UC16dBP
UC16dBP
UC16dBP
UC16dBP
UC16dBP
UC16dBP
UC16dBP
UC16dBP
UC16dBP
UC16dBP
UC16a-cBP
UC16a-cBP
UC16a-cBP
UC16a-cBP
UC16a-cBP
UC16a-cBP
UC16a-cBP
UC16a-cBP
UC16a-cBP
UC16a-cBP
UC16a-cBP
UC16a-cBP
UC16a-cBP
UC16a-cBP
UC16a-cBP
UC16a-cBP
UC15
UC15
UC15
UC15

Rørdal-1 corehole (R1)
10276
14.87
10239
27.10
10180
49.00
10113
70.15
10071
81.00

Lower Maastrichtian
Lower Maastrichtian
Lower MaastrichtianCampanian/MaastrichtianUpper Campanian

UC19
UC19
UC19
UC16dBP
UC16a-cBP

Rørdal quarry outcrop (Rq)4617209.1
Upper Maastrichtian46171610
Upper Maastrichtian

UC20b-cBP
UC20b-cBP

Calcareous nannofossils...boreal Campanian-Maastrichtian...Denmark

pixels/μm (1 pixel = 0.033μm), with an accuracy of measurement of ±2 pixels (0.066μm).
4. Systematic palaeontology
The terminology used in the descriptions below follows theguidelines of Young et al. (1997). In the taxonomic descriptions, L = length, W = width, WCA = width of the central area, LCA = length of the central area, E = ellipticity(ratio between coccolith length and width), central area
opening (= mean ratio between LCA/L and WCA/W).
Recorded occurrences in the studied sections: FO = firstoccurrence, LO = last occurrence. The higher taxonomy
follows the classification of Bown & Young (1997). All references prior to 1998 can be found in Perch-Nielsen (1985)
and Bown (1998). Others are given in the reference list.
4.1. Heterococcoliths
Imbricating muroliths (loxoliths)
Order EIFFELLITHALES Rood et al., 1971
Family CHIASTOZYGACEAE Rood et al., 1973
emend. Varol & Girgis, 1994
Central-area axial cross or bars
Ahmuellerella octoradiata (Górka, 1957) Reinhardt, 1966
Pl.1, fig.1
Ahmuellerella regularis (Górka, 1957) Reinhardt &Górka, 1967
Pl.1, figs 2, 3
Bukrylithus ambiguus Black, 1971
Pl.1, figs 4, 5
Heteromarginatus bugensis (Górka, 1957) Crux in Cruxet al., 1982
Pl.1, figs 6, 7
Monomarginatus quaternarius Wind & Wise in Wise &Wind, 1977
Pl.1, fig.8
Staurolithites dicandidula (Bergen in Bralower &Bergen, 1998) comb. nov.
Pl.1, figs 9-11
Basionym: Vagalapilla dicandidula Bergen in Bralower &Bergen, 1998: p.76, pl.1, figs 1a, 1b (holotype). (Bralower,
T.J. & Bergen, J.A. 1998. Cenomanian-Santonian calcareous nannofossil biostratigraphy of a transect of cores drilled
across the western interior seaway. In: M.A. Arthur & W.E.
Dean (Eds). Stratigraphy and Paleoenvironments of theCretaceous Western Interior Seaway, USA. SEPM Concepts in Sedimentology and Paleontology, 6: 59–77.)
Staurolithites elongatus (Bukry, 1969) Burnett, 1997aPl.1, fig.12
Staurolithites flavus Burnett, 1997aPl.1, fig.13

41

Staurolithites imbricatus (Gartner, 1968) Burnett, 1997aPl.1, figs 14, 15
Staurolithites laffittei Caratini, 1963
Pl.1, fig.16
Staurolithites mielnicensis (Górka, 1957) Perch-Nielsen,
1968 sensu Crux in Lord, 1982
Pl.1, figs 17, 18
Staurolithites zoensis Burnett, 1997aPl.1, figs 19, 20
Central-area transverse bar
Gorkaea pseudanthophorus (Bramlette & Martini, 1964)
Varol & Girgis, 1994
Pl.1, figs 21, 22
Placozygus fibuliformis (Reinhardt, 1964) Hoffmann,
1970
Pl.1, figs 23-25
Reinhardtites anthophorus (Deflandre, 1959)
Perch-Nielsen, 1968
Pl.1, figs 26-28
Reinhardtites levis Prins & Sissingh in Sissingh, 1977
Pl.1, figs 29-31
Tranolithus minimus (Bukry, 1969) Perch-Nielsen, 1984
Pl.1, fig.32
Tranolithus orionatus (Reinhardt, 1966a) Perch-Nielsen,
1968
Pl.1, fig.33
Tranolithus stemmerikii Thibault & Sheldon sp. nov.
Pl.1, figs 34, 35, Pl.2, figs 1-3. Derivation of name: After
Lars Stemmerik, Head of the Department of Geography
and Geology at the University of Copenhagen, co-founderof the CRC group. Diagnosis: A medium-sized species ofTranolithus with a reduced and small transverse bar and astrongly birefringent inner cycle. The inner cycle appears
to extend into four plates that almost fill the central area.
Opposing plates go into extinction together when rotatedthrough 30˚ (Pl.1, figs 33, 34; Pl.2, figs 1, 2). The plates
covering the central area surround two openings, alignedwith the long axis of the coccolith (Pl.1, fig.33; Pl.2, fig.1).
The bar is apparent only in well-preserved specimens andonly through certain angles when rotated. Differentiation:
This new species is distinguished from other species ofTranolithus by its small transerve bar and by its strongly
birefringent inner cycle, composed of four plates that fill
the central area and surround two openings that are alignedwith the long axis. In recrystallised specimens, the central
area is entirely covered by the four plates (Pl.2, fig.3). This
species is documented and referred to as Zeugrhabdotus
praesigmoides in Sheldon (2008, Pl.1, figs 15-18).
Holotype: Pl.1, figs 33-35 (L = 6.8μm, W = 5μm).
Paratypes: Pl.2, figs 1-4. Type locality: Stevns-1 corehole,Denmark. Type level: S1-7076, Upper Campanian(UC16aBP-cBP). Occurrence: Stevns-1, Rørdal-1; Upper
Campanian–Campanian/Maastrichtian boundary interval(UC15eBP-UC16). Remarks: Both in Stevns-1 and Rørdal-Thibault

42

1, the LO of this species was recorded within UC16dBP. It
thus has potential as a biostratigraphic indicator of theCampanian/Maastrichtian boundary interval.
Zeugrhabdotus acanthus Reinhardt, 1965
Pl.2, figs 5-7
Zeugrhabdotus bicrescenticus (Stover, 1966) Burnett in
Gale et al., 1996
Pl.2, fig.8
Zeugrhabdotus blowii Lees, 2007
Pl.2, figs 9, 10
Zeugrhabdotus diplogrammus (Deflandre in Deflandre &Fert, 1954) Burnett in Gale et al., 1996
Pl.2, figs 11, 12
Zeugrhabdotus cf. Z. erectus (Deflandre in Deflandre &Fert, 1954) Reinhardt, 1965
Pl.2, fig.13
Zeugrhabdotus erectus (Deflandre in Deflandre & Fert,
1954) Reinhardt, 1965
Pl.2, fig.14
Zeugrhabdotus sigmoides (Bramlette & Sullivan, 1961)
Bown & Young, 1997
Pl.2, figs 15-17
Zeugrhabdotus trivectis Bergen, 1994
Pl.2, figs 18, 19
Central-area net or vacant

Family EIFFELLITHACEAE Reinhardt, 1965
Eiffellithus collis Hoffman, 1970
Pl.3, fig.2
Eiffellithus eximius (Stover, 1966) Perch-Nielsen, 1968
Pl.3, figs 3, 4
Eiffellithus gorkae Reinhardt, 1965
Pl.3, fig.5
Eiffellithus parallelus Perch-Nielsen, 1973
Pl.3, figs 6, 7
Eiffellithus turriseiffelii (Deflandre in Deflandre & Fert,
1954) Reinhardt, 1965
Pl.3, figs 8, 9
Helicolithus anceps (Górka, 1957) Noël, 1970
Pl.3, figs 10, 11
Helicolithus trabeculatus (Górka, 1957) Verbeek, 1977
Pl.3, fig.12
Family RHAGODISCACEAE Hay, 1977
Rhagodiscus angustus (Stradner, 1963) Reinhardt, 1971
Pl.3, figs 13, 14
Rhagodiscus plebeius Perch-Nielsen, 1968
Pl.3, fig.15
Rhagodiscus reniformis Perch-Nielsen, 1973
Pl.3, fig.16
Rhagodiscus splendens (Deflandre, 1953) Verbeek, 1977
Pl.3, fig.17

Loxolithus armilla (Black in Black & Barnes, 1959)
Noël, 1965
Pl.2, fig.20
Loxolithus thiersteinii (Roth, 1973) Lees & Bown, 2005
Pl.2, fig.21

Order STEPHANOLITHIALES Bown & Young, 1997

Central-area diagonal crossFamily STEPHANOLITHIACEAE Black, 1968

Chiastozygus amphipons (Bramlette & Martini, 1964)
Gartner, 1968
Pl.2, fig.22
Chiastozygus antiquus (Perch-Nielsen, 1973) Burnett,
1997aPl.2, figs 23-25
Chiastozygus bifarius Bukry, 1969
Pl.2, figs 26-28
Chiastozygus litterarius (Górka, 1957) Manivit, 1971
Pl.2, fig.29
Chiastozygus spissus Bergen in Bralower & Bergen,
1998
Pl.2, figs 30, 31

Corollithion completum Perch-Nielsen, 1973
Pl.3, fig.18
Corollithion exiguum Stradner, 1961
Pl.3, fig.19
Corollithion? madagaskarensis Perch-Nielsen, 1973
Pl.3, fig.20
Cylindralithus serratus Bramlette & Martini, 1964
Pl.3, fig.21
Cylindralithus nudus Bukry, 1969
Pl.3, fig.22
Cylindralithus? nieliae Burnett, 1997aPl.3, figs 23, 24
Perchnielsenella stradneri (Perch-Nielsen, 1973) Watkins
in Watkins & Bowdler, 1984
Pl.3, fig.25
Rhombolithion rhombicum (Stradner & Adamiker, 1966)
Black, 1973
Pl.3, fig.26
Rotelapillus Noël, 1973
Rotelapillus sp.
Pl.3, figs 27, 28

Central-area closed
Neocrepidolithus cf. N. cohenii (Perch-Nielsen, 1968)
Perch-Nielsen, 1984
Pl.2, figs 32-35
Neocrepidolithus ruegenensis Burnett, 1997aPl.3, fig.1

Non-imbricating muroliths(loxoliths and/or protoliths)

Calcareous nannofossils...boreal Campanian-Maastrichtian...DenmarkNon-imbricating (or radial) placoliths
and related taxa

43

Pl.4, figs 23-26
Family CRETARHABDACEAE Thierstein, 1973

Order PODORHABDALES Rood et al.
emend. Bown, 1987
Family AXOPODORHABDACEAE Bown & Young,1997
Cribrocorona gallica (Stradner, 1963) Perch-Nielsen,
1973
Pl.3, fig.29
Cribrosphaerella circula (Risatti, 1973) Lees, 2007
Pl.3, fig.30
Cribrosphaerella daniae Perch-Nielsen, 1973
Pl.3, figs 31, 32
Cribrosphaerella ehrenbergii (Arkhangelsky, 1912)
Deflandre in Piveteau, 1952
Pl.3, figs 33-35
Nephrolithus miniporus Reinhardt & Górka, 1967
Pl.4, figs 1, 5
Nephrolithus frequens Górka, 1957
Pl.4, figs 2-5
Perissocyclus fenestratus (Stover, 1966) Black, 1971
Pl.4, figs 6-10
Tetrapodorhabdus decorus (Deflandre in Deflandre &Fert, 1954) Wind & Wise in Wise & Wind, 1977
Pl.4, figs 11, 12

Cretarhabdus conicus Bramlette & Martini, 1964
Pl.4, fig.27
Grantarhabdus coronadventis (Reinhardt, 1966) Grün in
Grün & Allemann, 1975
Pl.4, figs 28, 29
Miravetesina bergenii Lees, 2007
Pl.4, fig.30
Retecapsa angustiforata Black, 1971
Pl.4, figs 31-33
Retecapsa crenulata (Bramlette & Martini, 1964) Grün in
Grün & Allemann, 1975
Pl.4, figs 34, 35; Pl.5, fig.1
Retecapsa ficula (Stover, 1966) Burnett, 1997aPl.5, figs 2, 3
Retecapsa schizobrachiata (Gartner, 1968) Grün in Grün
& Allemann, 1975
Pl.5, figs 4, 5
Retecapsa surirella (Deflandre in Deflandre & Fert,
1954) Grün in Grün & Allemann, 1975
Pl.5, figs 6-13. Remarks: A central cross is not routinely
observed in R. surirella with the LM in this material. In
some better-preserved specimens, the axial cross can be oserved by rotating and focusing up and down (Pl.5, figs 1013).

Family BISCUTACEAE Black, 1971
Family PREDISCOSPHAERACEAE Rood et al., 1971
Biscutum constans (Górka, 1957) Black in Black &Barnes, 1959 var. constans
Pl.4, figs 13, 14
Biscutum constans (Górka, 1957) Black in Black &Barnes, 1959 var. ellipticum (Górka, 1957)
Pl.4, fig.15
Remarks: B. constans and B. ellipticum are herein considered to be morphotypes, rather than distinct species, following the work of Bornemann & Mutterlose (2006), whofound the overall population of constans to have a smaller
range of sizes than ellipticum, but strongly overlapped withthe latter, and so size was determined not to be a useful criterion upon which to distinguish between the taxa; thesecan only be differentiated by the structure of the central
area - small in constans and large and perforated in ellipticum.
Biscutum coronum Wind & Wise in Wise & Wind, 1977
Pl.4, figs 16, 17
Biscutum magnum Wind & Wise in Wise & Wind, 1977
Pl.4, fig.18
Biscutum melaniae (Górka, 1957) Burnett, 1997aPl.4, figs 19, 20
Biscutum notaculum Wind & Wise in Wise & Wind, 1977
Pl.4, figs 21, 22
Discorhabdus ignotus (Górka, 1957) Perch-Nielsen, 1968

Prediscosphaera arkhangelskyi (Reinhardt, 1965) PerchNielsen, 1984
Pl.5, figs 14-17. Remarks: The occurrence of this species
in Stevns-1 and Rørdal-1 is consistent until the top ofUC16dBP, where it becomes more sporadic. The LO of thisrare species was recorded within UC20aBP in Rørdal-1 andat the base of UC20bBP-cBP in Stevns-1. Perch-Nielsen(1979) noted its LO within CC23b (equivalent to UC17 of
Burnett, 1998).
Prediscosphaera stoveri (Perch-Nielsen, 1968) Shafik &Stradner, 1971
Pl.5, figs 18, 19
Prediscosphaera borealensis sp. nov.
Pl.5, figs 20-25. Derivation of name: Referring to thenorthern high latitudes, from where this species is described. Diagnosis: A small, elliptical species of Prediscosphaera with a thin, axial cross, a thin outer cycle, and awide inner cycle, which tends to enclose the central area
(Pl.4, figs 20, 21). The inner cycle exhibits an anticlockwise spiral interference pattern and almost fills the central
area around the axial cross. The axial cross appears veryslightly offset from the axes of the coccolith, and its arms
are slightly sigmoidal. Only a few better-preserved speci-

Thibault

44

Width

Ellipticity

Length central area (LCA)

Width central area (WCA)

Ratio LCA/length

Ratio WCA/width

Central area opening

Outer cycle width

P. borealensis (holotype)P. borealensis
P. borealensis
P. borealensis
P. borealensis
P. borealensis
P. borealensis
P. borealensis
P. borealensis
P. borealensis
P. borealensis
P. borealensis
P. borealensis
P. borealensis
P. borealensis
Mean
Maximum
Minimum

5.10
5.00
5.20
3.80
4.17
3.99
3.99
4.07
4.15
4.01
3.79
3.86
3.87
4.85
5.30
4.34
5.30
3.79

3.60
3.40
3.60
2.63
3.26
2.95
2.54
3.17
2.91
2.88
2.64
2.94
3.13
3.40
3.50
3.10
3.60
2.54

1.42
1.47
1.44
1.44
1.28
1.35
1.57
1.28
1.43
1.39
1.44
1.31
1.24
1.43
1.51
1.40
1.57
1.24

1.70
1.90
1.80
1.15
1.05
1.27
1.39
1.16
0.98
0.85
0.78
0.93
0.90
1.40
1.70
1.26
1.90
0.78

1.30
1.30
1.30
0.77
1.20
0.93
0.65
0.89
0.74
0.55
0.55
0.64
0.52
0.90
1.00
0.88
1.30
0.52

0.33
0.38
0.35
0.30
0.25
0.32
0.35
0.29
0.24
0.21
0.21
0.24
0.23
0.29
0.32
0.29
0.38
0.21

0.36
0.38
0.36
0.29
0.37
0.32
0.26
0.28
0.25
0.19
0.21
0.22
0.17
0.26
0.29
0.28
0.38
0.17

0.35
0.38
0.35
0.30
0.31
0.32
0.30
0.28
0.25
0.20
0.21
0.23
0.20
0.28
0.30
0.28
0.38
0.20

0.50
0.60
0.60
0.50
0.50
0.50
0.50
0.60
0.60
0.60
0.60
0.60
0.70
0.60
0.60
0.57
0.70
0.50

4.24
4.20
3.41
4.89
3.74
3.74
3.52
3.78
5.20
4.59
4.15
4.00
3.50
4.30
4.00
Mean 4.08
Maximum 5.20
Minimum 3.41

3.21
3.13
2.52
3.62
3.59
3.05
2.65
2.42
4.00
3.71
3.02
3.40
2.80
3.60
3.20
3.19
4.00
2.42

1.32
1.34
1.35
1.35
1.04
1.23
1.33
1.56
1.30
1.24
1.37
1.18
1.25
1.19
1.25
1.29
1.56
1.04

1.89
1.60
1.45
2.36
1.50
1.92
1.37
1.82
2.20
2.13
1.89
1.60
1.40
2.00
1.50
1.78
2.36
1.37

1.18
1.31
0.88
1.57
1.29
1.25
1.03
0.95
1.44
1.56
1.41
0.80
1.00
1.20
0.90
1.18
1.57
0.80

0.45
0.38
0.43
0.48
0.40
0.51
0.39
0.48
0.42
0.46
0.46
0.40
0.40
0.47
0.38
0.43
0.51
0.38

0.37
0.42
0.35
0.43
0.36
0.41
0.39
0.39
0.36
0.42
0.47
0.24
0.36
0.33
0.28
0.37
0.47
0.24

0.41
0.40
0.39
0.46
0.38
0.46
0.39
0.44
0.39
0.44
0.46
0.32
0.38
0.40
0.33
0.40
0.46
0.32

0.80
0.60
0.60
0.80
0.90
0.70
0.60
0.80
0.70
1.00
0.80
0.90
0.60
0.80
0.80
0.76
1.00
0.60

SampleLengthwith Pl.5, figs 20-25), and the different composition of theaxial cross. In P. stoveri and P. spinosa, the arms of thecross are straight and aligned with the axes of the coccolith (Pl.5, figs 18, 19; Pl.6, figs 8-10), whilst in the new
species, the arms are slightly offset from the axes andslightly sigmoidal (Pl.5, figs 20, 21). The new species differs from P. arkhangelskyi (Pl.5, figs 14-16) by its smallersize, the more enclosed central area, and the slightly offset
and slightly sigmoidal axial cross. Holotype: Pl.5, fig.20 (L= 5.1μm, W = 3.6μm; see Table 2 for range of dimensions).

Speciesmens were found with the axial cross present in this study.
In most specimens encountered, the cross was absent (Pl.5,
figs 22-25). It is possible that the cross is not visible inproximal view, where the inner cycle virtually fills the central area (Pl.5, figs 24, 25). Differentiation: This species
exhibits the same size range as P. stoveri (Table 2, Figure2). It can be distinguished from the latter, and from P. spinosa (Pl.6, figs 8-10), by the spiral pattern exhibited by the
inner cycle, the relatively thinner outer cycle, the more enclosed central area (Figure 2, and compare Pl.5, figs 18, 19

7081
7081
7081
1131
1131
1131
1131
1131
1131
1131
1131
1131
1131
7076
7076

1131
1131
1131
1131
1131
1131
1131
1131
1131
1131
1131
1463
1463
1463
1463

P. stoveri
P. stoveri
P. stoveri
P. stoveri
P. stoveri
P. stoveri
P. stoveri
P. stoveri
P. stoveri
P. stoveri
P. stoveri
P. stoveri
P. stoveri
P. stoveri
P. stoveri

Table 2: Dimensions and other morphometric parameters for Prediscosphaera borealensis sp. nov. and P. stoveri (15 specimens each)Calcareous nannofossils...boreal Campanian-Maastrichtian...Denmark45

Imbricating placoliths
Order WATZNAUERIALES Bown, 1987
Family WATZNAUERIACEAE Rood et al., 1971

Figure 2: Cross-plot of outer-cycle width versus central-area opening for
Prediscosphaera borealensis and P. stoveri (15 specimens each)Paratypes: Pl.5, figs 21-25. Type locality: Stevns-1 corehole, Denmark. Type level: S1-7081, Upper Campanian(UC16aBP-cBP). Occurrence: Stevns-1, Rørdal-1; Upper
Campanian-Lower Maastrichtian (UC16aBP-UC19).
Prediscosphaera cretacea (Arkhangelsky, 1912) Gartner,
1968
Pl.5, figs 26-30
Prediscosphaera grandis Perch-Nielsen, 1979
Pl.5, fig.31
Prediscosphaera incohatus (Stover, 1966) Burnett, 1997aPl.5, figs 32, 33
Prediscosphaera majungae Perch-Nielsen, 1973
Pl.5, figs 34, 35
Prediscosphaera mgayae Lees, 2007
Pl.6, figs 1-3. Remarks: This species may have potential asa stratigraphic tool. Both in Stevns-1 and Rørdal-1, its FO
lies within UC16dBP and its LO was recorded at the same
stratigraphic level as that of Reinhardtites levis, whichmarks the top of UC18. Lees (2007), who described thisspecies, noted its occurrence within UC17 in Tanzania.
Prediscosphaera microrhabdulina Perch-Nielsen, 1973
Pl.6, figs 4, 5
Prediscosphaera ponticula (Bukry, 1969) Perch-Nielsen,
1984
Pl.6, figs 6, 7
Prediscosphaera spinosa (Bramlette & Martini, 1964)
Gartner, 1968
Pl.6, figs 8-10
Family TUBODISCACEAE Bown & Rutledge in Bown
& Young, 1997
Manivitella pemmatoida (Deflandre in Manivit, 1965)
Thierstein, 1971
Pl.6, fig.11

Cyclagelosphaera alta Perch-Nielsen, 1979
Pl.6, fig.12
Cyclagelosphaera margerelii Noël, 1965
Pl.6, figs 13, 14
Cyclagelosphaera reinhardtii (Perch-Nielsen, 1968)
Romein, 1977
Pl.6, fig.15
Cyclagelosphaera rotaclypeata Bukry, 1969
Pl.6, fig.16
Cyclagelosphaera cf. C. tubulata Grün & Zweili, 1980
Pl.6, fig.18
Watznaueria barnesiae (Black, 1959) Perch-Nielsen,
1968
Pl.6, figs 19, 20, 31
Watznaueria biporta Bukry, 1969
Pl.6, figs 21, 22
Watznaueria britannica (Stradner, 1963) Reinhardt, 1964
Pl.6, figs 23, 24. Remarks: This species has been assumed
to become extinct in the Lower Cenomanian (Burnett et al.,1998), although a few specimens were found in the Coniacian (Burnett in Burnett et al., 1998, fig.6.4). In this study,
very rare specimens were found in one sample from theCampanian–Maastrichtian interval (UC16dBP) and threesamples from the Upper Maastrichtian (UC20aBP and
UC20dBP).
Watznaueria manivitiae sensu lato Bukry, 1973
Pl.6, figs 25-27
Watznaueria cf. W. ovata Bukry, 1969
Pl.6, figs 28, 29
Watznaueria ovata Bukry, 1969
Pl.6, fig.30
Other placolith-like coccoliths
Family ARKHANGELSKIELLACEAE Bukry, 1969
emend. Bown & Hampton in Bown & Young, 1997
Arkhangelskiella confusa Burnett, 1997aPl.6, fig.32
Arkhangelskiella cymbiformis Vekshina, 1959
Pl.6, figs 33, 34
Arkhangelskiella maastrichtiensis Burnett, 1997aPl.6, fig.35; Pl.7, fig.1
Broinsonia enormis (Shumenko, 1968) Manivit, 1971
Pl.7, fig.2
Broinsonia parca constricta Hattner et al., 1980
Pl.7, fig.3
Broinsonia parca parca (Stradner, 1963) Bukry, 1969
Pl.7, fig.4

Thibault

46

Broinsonia signata (Noël, 1969) Noël, 1970
Pl.7, figs 5, 6

Deflandre, 1947
Pl.8, figs 3, 4

Family KAMPTNERIACEAE Bukry, 1969 emend.
Bown & Hampton in Bown & Young, 1997
Gartnerago segmentatum (Stover, 1966) Thierstein, 1974
Pl.7, figs 7, 8
Gartnerago nanum Thierstein, 1974
Pl.7, figs 9, 10
Kamptnerius magnificus Deflandre, 1959
Pl.7, fig.11

Family LAPIDEACASSACEAE Bown & Young, 1997
Lapideacassis Black, 1971
Lapideacassis sp.
Pl.8, fig.5. Remarks: Three specimens of Lapideacassis
were found in Stevns-1 sediments. All of them were broken
and lacked the tapering termination(s) which allow identification at the species level.

Placoliths of uncertain affinity

Family MICRORHABDULACEAE Deflandre, 1963

Markalius apertus Perch-Nielsen, 1979
Pl.7, fig.12
Markalius inversus (Deflandre in Deflandre & Fert,
1954) Bramlette & Martini, 1964
Pl.7, figs 13, 14
Repagulum parvidentatum? (Deflandre in Deflandre &Fert, 1954) Forchheimer, 1972
Pl.7, figs 19-21

Lithraphidites carniolensis Deflandre, 1963
Pl.8, fig.6
Lithraphidites quadratus Bramlette & Martini, 1964
Pl.8, fig.7
Microrhabdulus undosus Perch-Nielsen, 1973
Pl.8, figs 8-10
Microrhabdulus decoratus Deflandre, 1959
Pl.8, figs 11-15

Muroliths

Family POLYCYCLOLITHACEAE Forchheimer,
1972 emend. Varol, 1992

Tortolithus caistorensis Crux in Crux et al., 1982
Pl.7, figs 15, 16
Tortolithus hallii (Bukry, 1969) Crux in Crux et al., 1982
Pl.7, figs 17, 18
Coccolith sp.
Pl.7, figs 22-26
4.2. Holococcoliths
Family CALYPTROSPHAERACEAE Boudreaux &Hay, 1969
Acuturris scotus (Risatti, 1973) Wind & Wise in Wise &Wind, 1977
Pl.7, fig.27
Calculites paulus Lees, 2007
Pl.7, figs 28-30
Calculites obscurus (Deflandre, 1959) Prins & Sissinghin Sissingh, 1977
Pl.7, fig.29
Calculites cf. C. rosalyniae Lees, 2007
Pl.7, figs 32, 33
Octolithus multiplus (Perch-Nielsen, 1973) Romein, 1979
Pl.7, figs 34, 35
Lucianorhabdus cayeuxii Deflandre, 1959
Pl.8, figs 1, 2

Micula concava (Stradner in Martini & Stradner, 1960)
Verbeek, 1976
Pl.9, figs 16, 17
Micula cubiformis Forchheimer, 1972
Pl.9, fig.18
Micula decussata Vekshina, 1959
Pl.9, fig.19. Remarks: As stated by Perch-Nielsen (1985),
M. decussata should be used rather than its synonym
M. staurophora, because the latter was poorly describedfrom Miocene sediments into which it may have been reworked.
Micula murus (Martini, 1961) Bukry, 1973
Pl.9, fig.20. Remarks: M. murus is generally restricted totropical regions (Thierstein, 1981; Lees, 2002). It occurs
in a very few samples of the uppermost Maastrichtian of
Stevns-1 (upper part of UC20dBP).
Micula swastica Stradner & Steinmetz, 1984
Pl.9, figs 21-23
Uniplanarius gothicus (Deflandre, 1959) Hattner & Wise,
1980
Pl.9, fig.24
Quadrum svabenickae Burnett, 1997aPl.9, fig.25

4.3. Nannoliths
INCERTAE SEDIS
Family BRAARUDOSPHAERACEAE Deflandre, 1947
Braarudosphaera bigelowii (Gran & Braarud, 1935)

Ceratolithoides indiensis Burnett, 1997b
Pl.9, figs 26-28

Calcareous nannofossils...boreal Campanian-Maastrichtian...Denmark

Ceratolithoides self-trailiae Burnett, 1997b
Pl.9, figs 29, 30. Remarks: Ceratolithoides species are
generally restricted to tropical regions (Lees, 2002). A few
specimens occur in samples from the Campanian/Maastrichtian boundary interval (UC16dBP), close to theLower/Upper Maastrichtian boundary (UC20aBP), and in
the uppermost Maastrichtian (upper part of UC20dBP).
Biantholithus sparsus Bramlette & Martini, 1964
Pl.9, figs 31, 32
4.4. Calcareous dinocysts
Thoracosphaera Kamptner, 1927
Thoracosphaera sp.
Pl.9, fig.33
Acknowledgements
The author is grateful to the Danish Natural Science ResearchCouncil (FNU) and the Natural Science Faculty, University of
Copenhagen for funding. I warmly thank Dorthe Samuelsen, Annette Ryge and John Boserup from GEUS for their hearty welcome in the palynologisk lab and for technical support. Thanks toThomas Hansen who helped in translating Stenestad (2005). I
want to express all my gratitude to the reviewers: Matthew Hampton, Dr. Liam Gallagher (both Network Stratigraphic) and Dr.
Jean Self-Trail (USGS). It was a piece of work and your comments were very helpful. Many, many thanks to the Editor, Jackie
A. Lees (UCL). I owe you more than one drink.

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Calcareous nannofossils...boreal Campanian-Maastrichtian...Denmark

Plate 1
Chiastozygaceae

Ahm. octoradiata S1-1337 Ahmuellerella regularis S1-1337 (2), S1-1305 (3)Heteromarginatus bugensis S1-1011 (6), S1-7072 (7)Bukrylithus ambiguus S1-1066 (4), S1-7076 (5)Mon. quaternarius S1-7086 Staur. dicandidula S1-1417 (9), S1-8180 (10)Staur. dicandidula S1-8180 Staur. elongatus R1-10113 Staur. flavus S1-1141

Staurolithites imbricatus S1-1380 (same specimen)

Staur. laffitei S1-1170

Staurolithites zoensis S1-8157 (same specimen)

Staurolithites mielnicensis S1-1380

Gorkaea pseudanthophorus S1-1463

Placozygus fibuliformis S1-1337 (23, 24 same specimen), S1-1272 (25)Reinhardtites anthophorus S1-1164 (26), S1-1055 (27, 28)

Tran. minimus S1-1029

Tran. orionatus S1-1044

Reinhardtites levis S1-1175 (29), S1-1009 (30)

Tranolithus stemmerikii S1-7076 (holotype, same specimen)

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Thibault

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Plate 2
Chiastozygaceae

Zeug. acanthus S1-7370

Tranolithus stemmerikii S1-7076 (1, 2 same specimen), S1-1044 (3), S1-1131 (4)Zeugrhabdotus acanthus S1-1207 (same specimen)

Z. bicrescenticus S1-1044 Zeugrhabdotus blowii S1-1432 (8), S1-1442 (9)Zeug. diplogrammus S1-1044 (11), S1-1386 (12)Z. cf. Z. erectus S1-7398

Zeug. sigmoides S1-1164 (15, 16 same specimen)

Zeugrhabdotus trivectis S1-1272 (same specimen)

Lox. thiersteinii S1-1283

Chiastozygus antiquus S1-8157 (23), Rq-461716 (24, 25 same specimen)

Chi. amphipons S1-1364

Chiastozygus bifarius S1-1411 (26, 27 same specimen), S1-1337 (28)Chi. spissus S1-5235

Zeug. erectus S1-8180

Zeug. sigmoides S1-1164

Lox. armilla R1-10071

Chi. litterarius R1-10239 Chi. spissus S1-8157

Neocrepidolithus cf. N. cohenii S1-1240 (32, 33 same specimen), S1-5623 (34, 35 same specimen)

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Calcareous nannofossils...boreal Campanian-Maastrichtian...Denmark

Plate 3
Chiastozygaceae, Eiffellithaceae, Rhagodiscaceae, Stephanolithiaceae, Axopodorhabdaceae

Neo. ruegenensis S1-1305 Eiffellithus collis S1-1109 Eiffellithus eximius S1-1109

Eiffellithus parallelus S1-1471 (6), S1-1305 (7)Hel. anceps S1-1447

Eiffellithus turriseiffelii S1-1471 (8), S1-8157 (9)Hel. trabeculatus S1-5260 Rhagodiscus angustus S1-1380

Eiffellithus gorkae S1-1337

Helicolithus anceps S1-1337

Rhag. plebeius S1-1004

Rhag. reniformis S1-7398 Rhag. splendens S1-1007

Cor. completum Rq-461720 Cor. exiguum Rq-461720 C.? madagaskarensis S1-1390

Cyl. serratus S1-1287

Cylindralithus? nieliae S1-1287

Cyl. nudus S1-1287

Rho. rhombicum S1-8180 Rotelapillus sp. S1-7074 (same specimen)

Cribrosphaerella daniae S1-1475 (31), S1-1422 (32)Per. stradneri S1-1287

Cribrocor. gallica S1-1469 Cribrosph. circula S1-1262

Cribrosphaerella ehrenbergii S1-1337 (33, 34), S1-1240 (35)Thibault

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Plate 4
Axopodorhabdaceae, Biscutaceae, Cretarhabdaceae

Nephrolithus miniporus
S1-5620 (1), S1-1471 (5)Nephrolithus frequens S1-1386 (2, 4), S1-1471 (3, 5)Perissocyclus fenestratus S1-1203 (6-8 same specimen and 9, 10 same specimen)

Tetrapodorhabdus decorus S1-1175 (11), S1-5620 (12) Biscutum constans var. constans S1-5209

Biscutum coronum S1-7084 (16), S1-7076 (17)Bisc. magnum S1-7076

Biscutum notaculum S1-1119 (same specimen)

Discorhabdus ignotus S1-8147

Disc. ignotus S1-8147

Grantarhabdus coronadventis R1-10276

Cret. conicus S1-1186

Retecapsa angustiforata S1-1114 (31), S1-1109 (32), S1-1087 (33)

B. constans var. ellipticum
S1-1066

Biscutum melaniae S1-1337 (19), S1-1170 (20)Mira. bergenii S1-1131

Retecapsa crenulata S1-1083 (34), S1-1044 (35)53

Calcareous nannofossils...boreal Campanian-Maastrichtian...Denmark

Plate 5
Cretarhabdaceae, Prediscosphaeraceae

Ret. crenulata S1-1186

Retecapsa ficula S1-1186 (same specimen)

Retecapsa schizobrachiata S1-1337

Retecapsa surirella S1-1262 (6), S1-1044 (7, 8 same specimen), S1-1170 (9), R1-10180 (10-13 same specimen)

Retecapsa surirella (see above)

Pre. arkhangelskyi S1-1044 (16), S1-1153 (17)Pre. arkhangelskyi S1-1207 (same specimen)

Prediscosphaera stoveri S1-1131 (18), S1-7076 (19)Pre. borealensis S1-7081

Prediscosphaera borealensis S1-7081 (21), S1-7076 (22), S1-1131 (23, 24), S1-1029 (25)

Prediscosphaera cretacea S1-1119 (26), S1-1474 (27), S1-1447 (28), S1-1175 (29), S1-1175 (30, side view)Pre. grandis S1-1175

Prediscosphaera incohatus S1-5253 (same specimen)

Pre. majungae S1-1473 (34), S1-1175 (35, side view)Thibault

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Plate 6Prediscosphaeraceae, Tubodiscaceae, Watznaueriaceae, Arkhangelskiellaceae

Prediscosphaera mgayae S1-1029 (1), S1-1175 (2, 3 same specimen)

Pre. microrhabdulina S1-1337 (4), S1-1337 (5, spine)

Pre. ponticula S1-1175 (notice node on 6)

Prediscosphaera spinosa S1-1422 (8, 10), S1-1442 (9)Man. pemmatoida S1-1337 Cyc. alta S1-1337

Cyc. margerelii S1-1011 (13), S1-1337 (14)Cyc. rotaclypeata S1-1337 Cyc. or Watz. coccosphereS1-1337

C. cf. C. tubulata S1-1337 Watznaueria barnesiae
S1-1044 (19, smaller form), S1-1044 (20, larger form)

Watznaueria biporta S1-1337 (same specimen)

Watznaueria britannica S1-1337

W. manivitiae s.l. S1-1337

Watz. manivitiae s.l. S1-1337 (same specimen)

Watznaueria cf. W. ovata S1-1337

Watz. ovata S1-1337

W. barnesiae coccosphere Ark. confusa S1-1175
S1-1011

Ark. cymbiformis S1-1305 (33), S1-1442 (34)

A. maastrichtiensis S1-1411

Cyc. reinhardtii S1-1337

Calcareous nannofossils...boreal Campanian-Maastrichtian...Denmark

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Plate 7
Arkhangelskiellaceae, Kamptneriaceae, incertae sedis heterococcoliths, holococcoliths

A. maastrichtiensis S1-1442 Broin. enormis S1-1066

B. parca constricta S1-1467 B. parca parca S1-1004

Gartnerago segmentatum S1-1272 (7), S1-1131 (8)Kam. magnificus S1-1153 Mark. apertus S1-5235

Gartnerago nanum S1-1066 (9), S1-1109 (10)Markalius inversus S1-1473 (13), S1-1462 (14)Tort. caistorensis S1-1066 Tortolithus hallii S1-1044 (17), S1-1066 (18)

B. signata S1-1017(5, 6 same specimen)

Tort. caistorensis S1-1066

R. parvidentatum? S1-1141 (19-21 same specimen)

Heterococcolith sp. S1-1447 (22-25 same specimen)

Het. sp. S1-1447

Acuturris scotus S1-1044

Calc. obscurus S1-1083

Calculites cf. C. rosalyniae S1-5235 (32), S1-1462 (33) Octolithus multiplus S1-1370 (34), S1-1044 (35)Calculites paulus S1-1141 (28, 29 same specimen)

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Plate 8
Holococcoliths, nannoliths

Lucianorhabdus cayeuxii S1-1044 (1), S1-1017 (2)Braarudosphaera bigelowii S1-7070 (3), S1-1364 (4) Lapideacassis sp. S1-1175

Lith. carniolensis S1-1207 Lith. quadratus S1-1386

M. undosus S1-1272 (8)Microrhabdulus decoratus S1-1272 (11-13 same specimen)

Microrhabdulus undosus
S1-1386 (9, 10 same specimen)

M. decoratus S1-1272 (14, 15 same specimen)

Micula concava S1-1447 (16, 17 same specimen)

Micula decussata S1-1300 Micula murus S1-1466

Micula swastica S1-1175 (21), S1-1283 (22, 23)

Uni. gothicus S1-1083

Ceratolithoides indiensis S1-1364 (26), S1-1310 (27, 28 same specimen)

Q. svabenickae S1-1004

Cer. self-trailiae Rq-461727 (29), S1-1353 (30)

Biantholithus sparsus S1-1066 (31), S1-5256 (32)

Micula cubiformis S1-1337

Thoracosphaera sp.
fragments S1-1066